Assessing the effect of prevalence on the predictive performance of species distribution models using simulated data

Aim The proportion of sampled sites where a species is present is known as prevalence. Empirical studies have shown that prevalence can affect the predictive performance of species distribution models. This paper uses simulated species data to examine how prevalence and the form of species environmental dependence affect the assessment of the predictive performance of models. Methods Simulated species data were based on various functions of simulated environmental data with differing degrees of spatial correlation. Seven model performance measures – sensitivity, specificity, class‐average (CA), overall prediction success, kappa (κ), normalized mutual information (NMI) and area under the receiver operating characteristic curve (AUC) – were applied to species models fitted by three regression methods. The response of the performance measures to prevalence was then assessed. Three probability threshold selection methods used to convert fitted logistic model values to presence or absence were also assessed. Results The study shows that the extent to which prevalence affects model performance depends on the modelling technique and its degree of success in capturing dominant environmental determinants. It also depends on the statistic used to measure model performance and the probability threshold method. The response based on κ generally preferred models with medium prevalence. All performance measures were least affected by prevalence when the probability threshold was chosen to maximize predictive performance or was based directly on prevalence. In these cases, the responses based on AUC, CA and NMI generally preferred models with small or large prevalence. Main conclusions The effect of prevalence on the predictive performance of species distribution models has a methodological basis. Relevant factors include the success of the fitted distribution model in capturing the dominant environmental determinant, the model performance measure and the probability threshold selection method. The fixed probability threshold method yields a marked response of model performance to prevalence and is therefore not recommended. The study explains previous empirical results obtained with real data.     

Minimum required number of specimen records to develop accurate species distribution models

Species distribution models (SDMs) are widely used to predict the occurrence of species. Because SDMs generally use presence‐only data, validation of the predicted distribution and assessing model accuracy is challenging. Model performance depends on both sample size and species’ prevalence, being the fraction of the study area occupied by the species. Here, we present a novel method using simulated species to identify the minimum number of records required to generate accurate SDMs for taxa of different pre‐defined prevalence classes. We quantified model performance as a function of sample size and prevalence and found model performance to increase with increasing sample size under constant prevalence, and to decrease with increasing prevalence under constant sample size. The area under the curve (AUC) is commonly used as a measure of model performance. However, when applied to presence‐only data it is prevalence‐dependent and hence not an accurate performance index. Testing the AUC of an SDM for significant deviation from random performance provides a good alternative. We assessed the minimum number of records required to obtain good model performance for species of different prevalence classes in a virtual study area and in a real African study area. The lower limit depends on the species’ prevalence with absolute minimum sample sizes as low as 3 for narrow‐ranged and 13 for widespread species for our virtual study area which represents an ideal, balanced, orthogonal world. The lower limit of 3, however, is flawed by statistical artefacts related to modelling species with a prevalence below 0.1. In our African study area lower limits are higher, ranging from 14 for narrow‐ranged to 25 for widespread species. We advocate identifying the minimum sample size for any species distribution modelling by applying the novel method presented here, which is applicable to any taxonomic clade or group, study area or climate scenario.     

Plotting receiver operating characteristic and precision–recall curves from presence and background data

1. The receiver operating characteristic (ROC) and precision–recall (PR) plots have been widely used to evaluate the performance of species distribution models. Plotting the ROC/PR curves requires a traditional test set with both presence and absence data (namely PA approach), but species absence data are usually not available in reality. Plotting the ROC/PR curves from presence‐only data while treating background data as pseudo absence data (namely PO approach) may provide misleading results.
2. In this study, we propose a new approach to calibrate the ROC/PR curves from presence and background data with user‐provided information on a constant c, namely PB approach. Here, c defines the probability that species occurrence is detected (labeled), and an estimate of c can also be derived from the PB‐based ROC/PR plots given that a model with good ability of discrimination is available. We used five virtual species and a real aerial photography to test the effectiveness of the proposed PB‐based ROC/PR plots. Different models (or classifiers) were trained from presence and background data with various sample sizes. The ROC/PR curves plotted by PA approach were used to benchmark the curves plotted by PO and PB approaches.
3. Experimental results show that the curves and areas under curves by PB approach are more similar to that by PA approach as compared with PO approach. The PB‐based ROC/PR plots also provide highly accurate estimations of c in our experiment.
4. We conclude that the proposed PB‐based ROC/PR plots can provide valuable complements to the existing model assessment methods, and they also provide an additional way to estimate the constant c (or species prevalence) from presence and background data.

     

Intra‐specific variability and plasticity influence potential tree species distributions under climate change

Aim To assess the effect of local adaptation and phenotypic plasticity on the potential distribution of species under future climate changes. Trees may be adapted to specific climatic conditions; however, species range predictions have classically been assessed by species distribution models (SDMs) that do not account for intra‐specific genetic variability and phenotypic plasticity, because SDMs rely on the assumption that species respond homogeneously to climate change across their range, i.e. a species is equally adapted throughout its range, and all species are equally plastic. These assumptions could cause SDMs to exaggerate or underestimate species at risk under future climate change. Location The Iberian Peninsula. Methods Species distributions are predicted by integrating experimental data and modelling techniques. We incorporate plasticity and local adaptation into a SDM by calibrating models of tree survivorship with adaptive traits in provenance trials. Phenotypic plasticity was incorporated by calibrating our model with a climatic index that provides a measure of the differences between sites and provenances. Results We present a new modelling approach that is easy to implement and makes use of existing tree provenance trials to predict species distribution models under global warming. Our results indicate that the incorporation of intra‐population genetic diversity and phenotypic plasticity in SDMs significantly altered their outcome. In comparing species range predictions, the decrease in area occupancy under global warming conditions is smaller when considering our survival–adaptation model than that predicted by a ‘classical SDM’ calibrated with presence–absence data. These differences in survivorship are due to both local adaptation and plasticity. Differences due to the use of experimental data in the model calibration are also expressed in our results: we incorporate a null model that uses survival data from all provenances together. This model always predicts less reduction in area occupancy for both species than the SDM calibrated with presence–absence. Main conclusions We reaffirm the importance of considering adaptive traits when predicting species distributions and avoiding the use of occurrence data as a predictive variable. In light of these recommendations, we advise that existing predictions of future species distributions and their component populations must be reconsidered.     

Many unreported crop pests and pathogens are probably already present

Invasive species threaten global biodiversity, food security and ecosystem function. Such incursions present challenges to agriculture where invasive species cause significant crop damage and require major economic investment to control production losses. Pest risk analysis (PRA) is key to prioritize agricultural biosecurity efforts, but is hampered by incomplete knowledge of current crop pest and pathogen distributions. Here, we develop predictive models of current pest distributions and test these models using new observations at subnational resolution. We apply generalized linear models (GLM) to estimate presence probabilities for 1,739 crop pests in the CABI pest distribution database. We test model predictions for 100 unobserved pest occurrences in the People's Republic of China (PRC), against observations of these pests abstracted from the Chinese literature. This resource has hitherto been omitted from databases on global pest distributions. Finally, we predict occurrences of all unobserved pests globally. Presence probability increases with host presence, presence in neighbouring regions, per capita GDP and global prevalence. Presence probability decreases with mean distance from coast and known host number per pest. The models are good predictors of pest presence in provinces of the PRC, with area under the ROC curve (AUC) values of 0.75–0.76. Large numbers of currently unobserved, but probably present pests (defined here as unreported pests with a predicted presence probability >0.75), are predicted in China, India, southern Brazil and some countries of the former USSR. We show that GLMs can predict presences of pseudoabsent pests at subnational resolution. The Chinese literature has been largely inaccessible to Western academia but contains important information that can support PRA. Prior studies have often assumed that unreported pests in a global distribution database represent a true absence. Our analysis provides a method for quantifying pseudoabsences to enable improved PRA and species distribution modelling.     

Assessing spatial distribution of Coffea arabica L. in Ethiopia's highlands using species distribution models and geospatial analysis methods

Though there is an increase in popularity of predictive modelling for assessing the geographical distribution of species, there is still a clear gap on explaining geospatial methods to derive the presence/absence of species in terms of geospatial extent besides the ambiguity of robust models. In this paper, we evaluate four major species distribution modelling methods: Artificial Neural Network (ANN), Support Vector Machines (SVM), Maximum Entropy (MaxEnt) and Generalized Linear Model (GLM) with pseudo absence and background absence data. To investigate the efficacy of these models, we present a case study using Coffea arabica L. species in Ethiopia as there was no species distribution modelling that has been done at a local scale especially in the coffee growing areas. We made predictions on 75% subsets and validation on 25% of the 112 presence of the species records that were collected from field observation and 0.5 m spatial resolution of true colour aerial photographs. Twelve biophysical explanatory variables; climatic, remote sensing based and landscape variables were employed in modelling. The results show that MaxEnt with pseudo absence data and SVM with background absence have highest area of understory coffee presence prediction with 12.2% and 23.1% area coverage of indigenous forest, respectively. The result from the model performance test using True Positive Rate (TPR) shows that GLM and SVM with pseudo absence data performed highest (TPR = 0.821). MaxEnt and SVM were the robust modelling methods (TPR = 0.964) using background absence data.     

Finessing atlas data for species distribution models

Aim The spatial resolution of species atlases and therefore resulting model predictions are often too coarse for local applications. Collecting distribution data at a finer resolution for large numbers of species requires a comprehensive sampling effort, making it impractical and expensive. This study outlines the incorporation of existing knowledge into a conventional approach to predict the distribution of Bonelli’s eagle (Aquila fasciata) at a resolution 100 times finer than available atlas data. Location Malaga province, Andalusia, southern Spain. Methods A Bayesian expert system was proposed to utilize the knowledge from distribution models to yield the probability of a species being recorded at a finer resolution (1 × 1 km) than the original atlas data (10 × 10 km). The recorded probability was then used as a weight vector to generate a sampling scheme from the species atlas to enhance the accuracy of the modelling procedure. The maximum entropy for species distribution modelling (MaxEnt) was used as the species distribution model. A comparison was made between the results of the MaxEnt using the enhanced and, the random sampling scheme, based on four groups of environmental variables: topographic, climatic, biological and anthropogenic. Results The models with the sampling scheme enhanced by an expert system had a higher discriminative capacity than the baseline models. The downscaled (i.e. finer scale) species distribution maps using a hybrid MaxEnt/expert system approach were more specific to the nest locations and were more contrasted than those of the baseline model. Main conclusions The proposed method is a feasible substitute for comprehensive field work. The approach developed in this study is applicable for predicting the distribution of Bonelli’s eagle at a local scale from a national‐level occurrence data set; however, the usefulness of this approach may be limited to well‐known species.     

A New Method for Estimation of Resource Selection Probability Function

ABSTRACT Weighted distributions can be used to fit various forms of resource selection probability functions (RSPF) under the use-versus-available study design (Lele and Keim 2006). Although valid, the numerical maximization procedure used by Lele and Keim (2006) is unstable because of the inherent roughness of the Monte Carlo likelihood function. We used a combination of the methods of partial likelihood and data cloning to obtain maximum likelihood estimators of the RSPF in a numerically stable fashion. We demonstrated the methodology using simulated data sets generated under the log—log RSPF model and a reanalysis of telemetry data presented in Lele and Keim (2006) using the logistic RSPF model. The new method for estimation of RSPF can be used to understand differential selection of resources by animals, an essential component of studies in conservation biology, wildlife management, and applied ecology.     

A fine‐grained spatial prediction model for the red‐listed vascular plant Scorzonera humilis

We explored the applied use of distribution modelling as a tool for making spatial predictions of occurrences of the red‐listed vascular plant species Scorzonera humilis in a study area in southeast Norway. Scorzonera is typical of extensively managed semi‐natural grasslands. A Maxent model was trained on all known records of the species, accurately georeferenced and gridded to fine resolution (grid cells of 25×25 m). Model performance was assessed on the training data by data‐splitting (by which some records were set off for evaluation) and on independent evaluation data collected in the field. Of the eight predictor variables used in the modelling, distance to roads and to arable land were most important followed by land‐cover class and altitude. Judged from the area under curve (AUC), the model was good to excellent and a significant, positive relationship was found between relative probabilities of occurrence predicted by the model and true probability of presence provided by the independently collected evaluation data. The model was used together with the evaluation data to estimate presence of Scorzonera humilis in 0.7% of the grid cells in the study area. The grid cells in which the model predicted highest probability for Scorzonera to be present had a true probability of presence of ca 12%, i.e. 17×higher than in an average cell. The present study demonstrates that, even when only simple predictor variables are available, spatial prediction modelling contributes important knowledge about rare species such as prevalence estimates, spatial prediction maps and insights into the species’ autecology. Spatial prediction modelling also makes cost‐efficient monitoring of rare species possible. However, it is pointed out that these benefits require evaluation of the model on independently sampled evaluation data.     

综述地方知识的生态应用价值

地方知识在解决生态问题上的价值在国外已经受到相当的重视,但在中国尚刚刚起步。通过综述国内外地方知识的相关文献,发现有关地方知识的研究最近几年呈逐渐上升的趋势,国外研究明显多于国内且比国内深入。整体而言,地方知识在生态应用中的价值可以体现在4个方面:作为背景知识、提供基础数据、提供未来方案以及参与生态管理。国内有关地方知识的研究还流于描述层面,大多在描述背景知识。而国外的研究则更多得聚焦于怎样利用地方知识来提供数据、提供新的规划及技术方法,并强化地方知识在生态管理中的作用。结合文献归纳了地方知识在生态应用梯度各等级中的具体作用,提出了一个地方知识的生态应用框架,以期指导中国的相关科研和实践工作。以国外的研究为鉴,地方知识在中国的生态应用未来具有很大空间,地方知识作为普遍知识的有效补充,应该充分参与数据、决策以及管理的全过程,提出地方知识在中国未来应用发展的几个重点突破的难点,期望地方知识在科学规划、资源管理中发挥其应有的价值。     

The effect of sample size on the accuracy of species distribution models: considering both presences and pseudo‐absences or background sites

Most high‐performing species distribution modelling techniques require both presences, and either absences or pseudo‐absences or background points. In this paper, we explore the effect of sample size, towards developing improved strategies for modelling. We generated 1800 virtual species with three levels of prevalence using ten modelling techniques, while varying the number of training presences (NTP) and the number of random points (NRP representing pseudo‐absences or background sites). For five of the ten modelling techniques we built two versions of models: one with an equal total weight (ETW) setting where the total weight for pseudo‐absence is equivalent to the total weight for presence, and another with an unequal total weight (UTW) setting where the total weight for pseudo‐absence is not required to be equal to the total weight for presence. We compared two strategies for NRP: a small multiplier strategy (i.e. setting NRP at a few times as large as NTP), and a large number strategy (i.e. using numerous random points). We produced ensemble models (by averaging the predictions from 30 models built with the same set of training presences and different sets of random points in equivalent numbers) for three NTP magnitudes and two NRP strategies. We found that model accuracy altered as NRP increased with four distinct patterns of performance: increasing, decreasing, arch‐shaped and horizontal. In most cases ETW improved model performance. Ensemble models had higher accuracy than the corresponding single models, and this improvement was pronounced when NTP was low. We conclude that a large NRP is not always an appropriate strategy. The best choice for NRP will depend on the modelling techniques used, species prevalence and NTP. We recommend building ensemble models instead of single models, using the small multiplier strategy for NRP with ETW, especially when only a small number of species presence records are available.     

The effect of a gradual response to the environment on species distribution modeling performance

Species distribution models (SDMs) have been widely used in ecology, biogeography, and conservation. Although ecological theory predicts that species occupancy is dynamic, the outputs of SDMs are generally converted into a single occurrence map, and model performance is evaluated in terms of success to predict presences and absences. The aim of this study was to characterize the effects of a gradual response in species occupancy to environmental gradients into the performance of SDMs. First we outline guidelines for the appropriate simulation of artificial species that allows controlling for gradualism and prevalence in the occupancy patterns over an environmental gradient. Second, we derive theoretical expected values for success measures based on presence‐absence predictions (AUC, Kappa, sensitivity and specificity). And finally we used artificial species to exemplify and test the effect of a gradual probabilistic occupancy response to environmental gradients on SDM performance. Our results show that when a species responds gradually to an environmental gradient, conventional measures of SDM predictive success based on presence‐absence cannot be expected to attain currently accepted performance values considered as good, even for a model that recovers perfectly well the true probability of occurrence. A gradual response imposes a theoretical expected value for these measures of performance that can be calculated from the species properties. However, irrespective of the statistical modeling strategy used and of how gradual the species response is, one can recover the true probability of occurrence as a function of environmental variables provided that species and sample prevalence are similar. Therefore, model performance based on presence‐absence should be judged against the theoretical expected value rather than to absolute values currently in use such as AUC > 0.8. Overall, we advocate for a wider use of the probability of occurrence and emphasize the need for further technical developments in this sense.     

An assessment of the effect of data partitioning on the performance of modelling algorithms for habitat suitability for ticks

A comparison of the performance of five modelling methods using presence/absence (generalized additive models, discriminant analysis) or presence-only (genetic algorithm for rule-set prediction, ecological niche factor analysis, Gower distance) data for modelling the distribution of the tick species Boophilus decoloratus (Koch, 1844) (Acarina: Ixodidae) at a continental scale (Africa) using climate data was conducted. This work explicitly addressed the usefulness of clustering using the normalized difference vegetation index (NDVI) to split original records and build partial models for each region (cluster) as a method of improving model performance. Models without clustering have a consistently lower performance (as measured by sensitivity and area under the curve [AUC]), although presence/absence models perform better than presence-only models. Two cluster-related variables, namely, prevalence (commonness of tick records in the cluster) and marginality (the relative position of the climate niche occupied by the tick in relation to that available in the cluster) greatly affect the performance of each model (P < 0.05). Both sensitivity and AUC are better for NDVI-derived clusters where the tick is more prevalent or its marginality is low. However, the total size of the cluster or its fragmentation (measured by Shannon's evenness index) did not affect the performance of models. Models derived separately for each cluster produced the best output but resulted in a patchy distribution of predicted occurrence. The use of such a method together with weighting procedures based on prevalence and marginality as derived from populations at each cluster produced a slightly lower predictive performance but a better estimation of the continental distribution of the tick. Therefore, cluster-derived models are able to effectively capture restricting conditions for different tick populations at a regional level. It is concluded that data partitioning is a powerful method with which to describe the climate niche of populations of a tick species, as adapted to local conditions. The use of this methodology greatly improves the performance of climate suitability models.     

Uncertainty of bioclimate envelope models based on the geographical distribution of species

Aim We explored the effects of prevalence, latitudinal range and spatial autocorrelation of species distribution patterns on the accuracy of bioclimate envelope models of butterflies. Location Finland, northern Europe. Methods The data of a national butterfly atlas survey (NAFI) carried out in 1991–2003 with a resolution of 10 × 10 km were used in the analyses. Generalized additive models (GAM) were constructed, for each of 98 species, to estimate the probability of occurrence as a function of climate variables. Model performance was measured using the area under the curve (AUC) of a receiver operating characteristic (ROC) plot. Observed differences in modelling accuracy among species were related to the species’ geographical attributes using multivariate GAM. Results Accuracies of the climate–butterfly models varied from low to very high (AUC values 0.59–0.99), with a mean of 0.79. The modelling performance was related negatively to the latitudinal range and prevalence, and positively to the spatial autocorrelation of the species distribution. These three factors accounted for 75.2% of the variation in the modelling accuracy. Species at the margin of their range or with low prevalence were better predicted than widespread species, and species with clumped distributions better than scattered dispersed species. Main conclusions The results from this study indicate that species’ geographical attributes highly influence the behaviour and uncertainty of species–climate models, which should be taken into account in biogeographical modelling studies and assessments of climate change impacts.     

A probabilistic approach to estimating species pools from large compositional matrices

Abstract. Species pools are increasingly recognized as important controls of local plant community structure and diversity. While existing approaches to estimate their content and size either rely on phytosociological expert knowledge or on simple response models across environmental gradients, the proposed application of phytosociological smoothing according to Beals exploits the full information of plant co‐occurrence patterns statistically. Where numerous representative compositional data are available, the new method yields robust estimates of the potential of sites to harbour plant species. To test the new method, a large phytosociological databank covering the forested regions of Oregon (US) was subsampled randomly and evenly across strata defined by geographic regions and elevation belts. The resulting matrix of species presence/absence in 874 plots was smoothed by calculating Beals' index of sociological favourability, which estimates the probability of encountering each species at each site from the actual plot composition and the pattern of species co‐occurrence in the matrix. In a second step, the resulting lists of sociologically probable species were intersected with complete species lists for each of 14 geographical subregions. Species pools were compared to observed species composition and richness. Species pool size exhibited much clearer spatial trends than plot richness and could be modelled much better as a function of climatic factors. In this framework the goal of modelling species pools is not to test a hypothesis, but to bridge the gap between manageable scales of empirical observation and the spatio‐temporal hierarchy of diversity patterns.     

Linking local ecological knowledge and habitat modelling to predict absolute species abundance on large scales

Assessing the spatial structure of abundance of a species is a basic requirement to carry out adequate conservation strategies. However, existing attempts to predict species abundance, particularly in absolute units and on large scales, are scarce and have led to weak results. In this work we present a scheme to obtain, in an affordable way, a predictive model of absolute animal abundance on large scales based on the modelling of data obtained from local ecological knowledge (LEK) and its calibration. To exemplify this scheme, we build and validate a predictive absolute abundance model of the endangered terrestrial tortoise Testudo graeca in Southeast Iberian Peninsula. For that purpose, we collected distribution and relative abundance data of T. graeca using a low cost methodology, such as LEK, by means of interviewing shepherds. The information from LEK was employed to build a predictive habitat-based model of relative abundance. The relative abundance model was transformed into an absolute abundance model by means of calibration with a classical absolute abundance sampling method such as distance sampling. The obtained absolute abundance model predicted the observed absolute abundances values well in independent locations when compared with other works (R ² = 36%) and thus can offer a cost-effective predictive ability. Our results show that reliable habitat-based predictive maps of absolute species abundance on regional scales can be obtained starting from low cost sampling methods of relative abundance, such as LEK, and its calibration.     

Can we model the probability of presence of species without absence data?

In ecological studies, it is useful to estimate the probability that a species occurs at given locations. The probability of presence can be modeled by traditional statistical methods, if both presence and absence data are available. However, the challenge is that most species records contain only presence data, without reliable absence data. Previous presence‐only methods can estimate a relative index of habitat suitability, but cannot estimate the actual probability of presence. In this study, we develop a presence and background learning algorithm (PBL) that is successful in modeling the conditional probability of presence of a simulated species. The model is trained by two completely separate sets: observed presence and background data. Assuming that the probability of presence is one for ‘prototypical presence’ locations where the habitats are maximally suitable for a species, we can estimate a constant that can calibrate the trained model into the actual probability of presence. Experimental results show that the PBL method performs similarly to a presence‐absence method, and significantly better than the widely used maximum entropy method. The new algorithm enables us to model the probability that a species occurs conditional on environmental covariates without absence data. Hence, it has potential to improve modeling of the geographical distributions of species.     

Bayesian nonparametric centered random effects models with variable selection

In a linear mixed effects model, it is common practice to assume that the random effects follow a parametric distribution such as a normal distribution with mean zero. However, in the case of variable selection, substantial violation of the normality assumption can potentially impact the subset selection and result in poor interpretation and even incorrect results. In nonparametric random effects models, the random effects generally have a nonzero mean, which causes an identifiability problem for the fixed effects that are paired with the random effects. In this article, we focus on a Bayesian method for variable selection. We characterize the subject‐specific random effects nonparametrically with a Dirichlet process and resolve the bias simultaneously. In particular, we propose flexible modeling of the conditional distribution of the random effects with changes across the predictor space. The approach is implemented using a stochastic search Gibbs sampler to identify subsets of fixed effects and random effects to be included in the model. Simulations are provided to evaluate and compare the performance of our approach to the existing ones. We then apply the new approach to a real data example, cross‐country and interlaboratory rodent uterotrophic bioassay.     

Why Would Plant Species Become Extinct Locally If Growing Conditions Improve?

Two assumptions underlie current models of the geographical ranges of perennial plant species: 1. current ranges are in equilibrium with the prevailing climate, and 2. changes are attributable to changes in macroclimatic factors, including tolerance of winter cold, the duration of the growing season, and water stress during the growing season, rather than to biotic interactions. These assumptions allow model parameters to be estimated from current species ranges. Deterioration of growing conditions due to climate change, e.g. more severe drought, will cause local extinction. However, for many plant species, the predicted climate change of higher minimum temperatures and longer growing seasons means, improved growing conditions. Biogeographical models may under some circumstances predict that a species will become locally extinct, despite improved growing conditions, because they are based on an assumption of equilibrium and this forces the species range to match the species-specific macroclimatic thresholds. We argue that such model predictions should be rejected unless there is evidence either that competition influences the position of the range margins or that a certain physiological mechanism associated with the apparent improvement in growing conditions negatively affects the species performance. We illustrate how a process-based vegetation model can be used to ascertain whether such a physiological cause exists. To avoid potential modelling errors of this type, we propose a method that constrains the scenario predictions of the envelope models by changing the geographical distribution of the dominant plant functional type. Consistent modelling results are very important for evaluating how changes in species areas affect local functional trait diversity and hence ecosystem functioning and resilience, and for inferring the implications for conservation management in the face of climate change.     

Hierarchical multi-scale occupancy estimation for monitoring wildlife populations

Occupancy estimation is an effective analytic framework, but requires repeated surveys of a sample unit to estimate the probability of detection. Detection rates can be estimated from spatially replicated rather than temporally replicated surveys, but this may violate the closure assumption and result in biased estimates of occupancy. We present a new application of a multi-scale occupancy model that permits the simultaneous use of presence–absence data collected at 2 spatial scales and uses a removal design to estimate the probability of detection. Occupancy at the small scale corresponds to local territory occupancy, whereas occupancy at the large scale corresponds to regional occupancy of the sample units. Small-scale occupancy also corresponds to a spatial availability or coverage parameter where a species may be unavailable for sampling at a fraction of the survey stations. We applied the multi-scale occupancy model to a hierarchical sample design for 2 bird species in the Black Hills National Forest: brown creeper (Certhia americana) and lark sparrow (Chondestes grammacus). Our application of the multi-scale occupancy model is particularly well suited for hierarchical sample designs, such as spatially replicated survey stations within sample units that are typical of avian monitoring programs. The model appropriately accounts for the non-independence of the spatially replicated survey stations, addresses the closure assumption for the spatially replicated survey stations, and is useful for decomposing the observation process into detection and availability parameters. This analytic approach is likely to be useful for monitoring at local and regional scales, modeling multi-scale habitat relationships, and estimating population state variables for rare species of conservation concern. © 2011 The Wildlife Society.